Metapopulation vicariance explains old endemics on young volcanic islands

Terrestrial plants and animals on oceanic islands occupy zones of volcanism found at intraplate localities and along island arcs at subduction zones. The organisms often survive as metapopulations, or populations of separate sub‐populations connected by dispersal. Although the individual islands and their local subpopulations are ephemeral and unstable, the ecosystem dynamism enables metapopulations to persist in a region, more or less in situ, for periods of up to tens of millions of years. As well as surviving on systems of young volcanic islands, metapopulations can also evolve there; tectonic changes can break up widespread insular metapopulations and produce endemics restricted to fewer islands or even a single island. These processes explain the presence of old endemic clades on young islands, which is often reported in molecular clock studies, and the many distribution patterns in island life that are spatially correlated with tectonic features. Metapopulations can be ruptured by sea floor subsidence, and this occurs with volcanic loading in zones of active volcanism and with sea floor cooling following its production at mid‐ocean ridges. Metapopulation vicariance will also result if an active zone of volcanism is rifted apart. This can be caused by the migration of an arc (by slab rollback) away from a continent or from another subduction zone, by the offset of an arc at transform faults and by sea floor spreading at mid‐ocean ridges. These mechanisms are illustrated with examples from islands in the Caribbean and the Pacific. Endemism on oceanic islands has usually been attributed to chance, long‐distance dispersal, but the processes discussed here will generate endemism on young volcanic islands by vicariance.     

Conflicting patterns of genetic and morphological variation in European Gentianella section Gentianella

Genetics in European Gentianella Moench sect. Gentianella were investigated using internal transcribed spacer (ITS) nrDNA, trn L-F cpDNA and Amplified Fragment Length Polymorphisms (AFLP) fingerprint data to analyse relationships within the section. Both a cladistic and a phenetic approach, as well as nonhierarchical analysis (PCA) and partitioning of molecular variance (AMOVA), were used. Sequence data did not resolve groups within the section. They reflected, however, geographical isolation between central European and eastern Carpathian ( G. lutescens and G. bulgarica ) as well as Caucasian samples ( G. caucasea ). AFLP data revealed a strong differentiation within G. germanica between populations of the Alps and north-western middle Europe, although there is hardly any morphological differentiation of taxonomic significance. Morphological similarities between G. aspera and G. pilosa were not reflected in genetics, and no support was found for G. aspera . Genetic similarities were not in line with morphology in G. lutescens and G. bulgarica . AFLP data only partly resolve relationships among taxa. High genetic similarities, as well as partitioning of molecular variance among taxa/populations of eastern middle Europe ( G. austriaca , G. bohemica , G. germanica , G. stiriaca ) and intermediate morphological characters in G. bohemica and G. stiriaca , might indicate reticulate evolution. The overall low sequence divergence within the section points to a very young origin of all taxa.  © 2005 The Linnean Society of London, Botanical Journal of the Linnean Society , 2005, 148 , 175–187.     

Population structure, genetic variation and morphological diversity in indigenous sheep of Ethiopia

We investigated genetic and morphological diversity and population structure of 14 traditional sheep populations originating from four ecological zones in Ethiopia (sub-alpine, wet highland, sub-humid lowland and arid lowland). All animals (n = 672) were genotyped for 17 microsatellite markers and scored for 12 morphological characters. The sheep were initially classified as fat-tailed (11 populations), thin-tailed (one population) and fat-rumped sheep (two populations). These classifications are thought to correspond to three consecutive introduction events of sheep from the Near-East into East Africa. For the 14 populations, allelic richness ranged from 5.87 to 7.51 and expected heterozygosity (H(E)) from 0.66 to 0.75. Genetic differentiations (F(ST) values) between all pairs of populations, except between sub-alpine populations, were significantly different from zero (P < 0.001). Cluster analysis of morphological characters and a dendrogram constructed from genetic distances were broadly consistent with the classification into fat-tailed, thin-tailed and fat-rumped sheep. Bayesian cluster analysis using microsatellite markers indicated that there has been further genetic differentiation after the initial introduction of sheep into Ethiopia. Investigation of factors associated with genetic variation showed that an isolation-by-distance model, independently of other factors, explained most of the observed genetic variation. We also obtained a strong indication of adaptive divergence in morphological characters, patterns of morphological variation being highly associated with ecology even when the effect of neutral genetic divergence (F(ST)) was parcelled out in partial Mantel tests. Using a combination of F(ST) values, Bayesian clustering analysis and morphological divergence, we propose a classification of Ethiopian sheep into six breed groups and nine breeds.     

Pleiotropic effects on mandibular morphology II: differential epistasis and genetic variation in morphological integration

The evolution of morphological modularity through the sequestration of pleiotropy to sets of functionally and developmentally related traits requires genetic variation in the relationships between traits. Genetic variation in relationships between traits can result from differential epistasis, where epistatic relationships for pairs of loci are different for different traits. This study maps relationship quantitative trait loci (QTLs), specifically QTLs that affect the relationship between individual mandibular traits and mandible length, across the genome in an F2 intercross of the LG/J and SM/J inbred mouse strains (N = 1045). We discovered 23 relationship QTLs scattered throughout the genome. All mandibular traits were involved in one or more relationship QTL. When multiple traits were affected at a relationship QTL, the traits tended to come from a developmentally restricted region of the mandible, either the muscular processes or the alveolus. About one-third of the relationship QTLs correspond to previously located trait QTLs affecting the same traits. These results comprise examples of genetic variation necessary for an evolutionary response to selection on the range of pleiotropic effects.     

Inter- and intraspecific genetic and morphological variation in a sibling pair of carabid species

Background

Pogonus littoralis and Pogonus chalceus are very close related species with quite different ecological preferences within salt marshes. We study the evolutionary processes in and between these presumably young species. Therefore, we compare the variation in ecologically relevant characters and the genetic variation within one of the species (intraspecific differentiation) with the variation of the two types of characters between the two species (interspecific variation). Data are compared between two independent sets of populations, one set at a small geographical scale (the ecologically diverse Guérande area in France) and the other set at a Atlantic-Mediterranean scale.

Results

Body and relative wing size and IDH1 allozyme data show that the intraspecific variation in P. chalceus is high and in the same range as the interspecific variation (P. chalceus versus P. littoralis). Based on neutral markers (other allozymes and mitochondrial DNA) on the other hand, the intraspecific variation in P. chalceus is much lower in comparison to the interspecific variation.

Conclusion

The different ecotypes in the highly polytypic species P. chalceus are as highly differentiated in ecological characters as true species, but are not recognised as such by screening neutral DNA polymorphisms. This can be interpreted as a case of ongoing speciation driven by natural selection adapting each ecotype to its respective ecological niche. The same ecological process can be recognised in the differentiation between the two sister species, where en plus reproductive isolation between the two gene pools occurred, allowing independent drift and mutation accumulation in neutral genetic characters.     

Riverscape genetic variation,migration patterns,and morphological variation of the threatened Round Rocksnail,Leptoxis ampla

Within riverine systems, headwater populations are hypothesized to harbour higher amounts of genetic distinctiveness than populations in the main stem of a river and display increased genetic diversity in large, downstream habitats. However, these hypotheses were mostly developed with insects and fish, and they have not been tested on many invertebrate lineages. Pleuroceridae gastropods are of particular ecological importance to rivers of eastern North America, sometimes comprising over 90% of macroinvertebrate biomass. Yet, virtually nothing is known of pleurocerid landscape genetics, including whether genetic diversity follows predictions made by hypotheses developed on more mobile species. Moreover, the commonly repeated hypothesis that intraspecific morphological variation in gastropods results from ecophenotypic plasticity has not been well tested on pleurocerids. Using 2bRAD‐seq to discover single nucleotide polymorphisms, we show that the threatened, Cahaba River endemic pleurocerid, Leptoxis ampla, has limited gene flow among populations and that migration is downstream‐biased, conflicting with previous hypotheses. Both tributary and main stem populations harbour unique genomic profiles, and genetic diversity was highest in downstream populations. Furthermore, L. ampla shell morphology was more correlated with genetic differences among individuals and populations than habitat characteristics. We anticipate similar genetic and demographic patterns to be seen in other pleurocerids, and hypotheses about gene flow and population demographics that were based on more mobile taxa often, but not always, apply to freshwater gastropods. From a conservation standpoint, genetic structure of L. ampla populations suggests distinctive genetic diversity is lost with localized extirpation, a phenomenon common across the range of Pleuroceridae.     

4种形态类型诸葛菜的等位酶变异及遗传多样性分析

以采自江苏省南京市4个样点的4种不同形态类型(紫花毛果、紫花光果、白花光果和白花毛果)诸葛菜〔Orychophragmus violaceus(L.)O.E.Schulz〕63个单株为研究对象,采用不连续聚丙烯酰胺凝胶电泳技术(SDS-PAGE)对过氧化物酶同工酶(POD)、乙醇脱氢酶(ADH)、谷氨酸脱氢酶(GDH)、超氧化物歧化酶(SOD)和多酚氧化酶(PPO)酶谱进行了分析,并基于5个等位酶的酶谱分析结果对4种形态类型诸葛菜的遗传多样性进行了研究。结果表明:5个等位酶系统共包含12个位点32个等位基因,其中POD-1、POD-4、POD-5、POD-6、ADH-1、PPO-1、SOD-1和SOD-2为多态性位点;除具有共有谱带外,不同类型诸葛菜还具有各自特有的酶谱特征,其中,GDH-2的b带和c带分别是紫花类型和白花类型的特有谱带。紫花毛果、紫花光果和白花毛果类型的多态位点百分率(PPL)均为58.33%,而白花光果的PPL为66.67%;紫花毛果、紫花光果、白花光果和白花毛果每个位点的平均等位基因数分别为2.42、2.25、2.42和1.83,平均期望杂合度分别为0.35、0.30、0.38和0.29。4种形态类型诸葛菜的总基因多样度平均值为0.61,类型内基因多样度平均值为0.49,明显大于类型间基因多样度平均值(0.12);类型间的基因分化系数平均值为0.195,表明总基因多样性的80.5%来源于类型内。紫花光果和白花毛果类型间的遗传一致度最低(0.724 4)且遗传距离最大(0.322 5),而2个白花类型间的遗传一致度最高(0.954 1)且遗传距离最小(0.047 1)。研究结果显示:诸葛菜的种内变异程度很大、遗传分化程度较高,但在各类型内具有较高的遗传相似性。     

The population genetic theory of hidden variation and genetic robustness

One of the most solid generalizations of transmission genetics is that the phenotypic variance of populations carrying a major mutation is increased relative to the wild type. At least some part of this higher variance is genetic and due to release of previously hidden variation. Similarly, stressful environments also lead to the expression of hidden variation. These two observations have been considered as evidence that the wild type has evolved robustness against genetic variation, i.e., genetic canalization. In this article we present a general model for the interaction of a major mutation or a novel environment with the additive genetic basis of a quantitative character under stabilizing selection. We introduce an approximation to the genetic variance in mutation-selection-drift balance that includes the previously used stochastic Gaussian and house-of-cards approximations as limiting cases. We then show that the release of hidden genetic variation is a generic property of models with epistasis or genotype-environment interaction, regardless of whether the wild-type genotype is canalized or not. As a consequence, the additive genetic variance increases upon a change in the environment or the genetic background even if the mutant character state is as robust as the wild-type character. Estimates show that this predicted increase can be considerable, in particular in large populations and if there are conditionally neutral alleles at the loci underlying the trait. A brief review of the relevant literature suggests that the assumptions of this model are likely to be generic for polygenic traits. We conclude that the release of hidden genetic variance due to a major mutation or environmental stress does not demonstrate canalization of the wild-type genotype.     

High intraspecific genetic and morphological variation in the pioneer lichen Cladonia rei colonising slag dumps

This study investigates the genetic and morphological variability of the lichen Cladonia rei inhabiting strongly contaminated postsmelting slag dumps in southern Poland. Altogether, 27 C. rei samples were analysed, including 17 from a single population in one dump. The phylogenetic analysis includes samples of C. rei, outgroup species, and external sequences of Cladonia section representatives from GenBank. Comparative analysis of the internal transcribed spacer (ITS) rDNA sequences revealed the presence of 19 C. rei haplotypes overall, including several of the most frequent, of which 11 are represented by single individuals only. As many as 12 haplotypes were recorded within a single population. Three strongly supported monophyletic clades comprised of specimens from different geographical regions were recovered. Morphometric analysis showed great phenotypic variability within particular clades. Apart from a full range of previously known morphological forms of the species, an additional specific morphotype was recognised in the dumps; however, its representatives do not create a monophyletic group. High genetic variability within a single population suggests that C. rei has a great potential for colonising anthropogenic habitats. This attribute emphasises the role of this lichen as an essential pioneer in the early stages of natural regeneration of such sites.     

Seasonal genetic variation associated with population dynamics of a poecilogonous polychaete worm

Poecilogonous species show variation in developmental mode, with larvae that differ both morphologically and ecologically. The spionid polychaete Pygospio elegans shows variation in developmental mode not only between populations, but also seasonally within populations. We investigated the consequences of this developmental polymorphism on the spatial and seasonal genetic structure of P. elegans at four sites in the Danish Isefjord‐Roskilde‐Fjord estuary at six time points, from March 2014 until February 2015. We found genetic differentiation between our sampling sites as well as seasonal differentiation at two of the sites. The seasonal genetic shift correlated with the appearance of new size cohorts in the populations. Additionally, we found that the genetic composition of reproductive individuals did not always reflect the genetic composition of the entire sample, indicating that variance in reproductive success among individuals is a likely explanation for the patterns of chaotic genetic patchiness observed during this and previous studies. The heterogeneous, unpredictable character of the estuary might maintain poecilogony in P. elegans as a bet‐hedging strategy in the Isefjord‐Roskilde‐Fjord complex in comparison with other sites where P. elegans are expected to be fixed to a certain mode of development.     

A contribution to the knowledge of myxomycetes diversity in volcanic islands

A list of myxomycetes collected in Pantelleria, a volcanic island located 110 km southwest of the island of Sicily (Italy), is presented with data on distribution. Forty-nine taxa were identified, 38 of which are new for Sicily. The myxomycete diversity of Pantelleria was also compared with that of other volcanic islands located in the Atlantic and Pacific oceans.     

The effect of epistasis on sexually antagonistic genetic variation

There is increasing evidence of segregating sexually antagonistic (SA) genetic variation for fitness in laboratory and wild populations, yet the conditions for the maintenance of such variation can be restrictive. Epistatic interactions between genes can contribute to the maintenance of genetic variance in fitness and we suggest that epistasis between SA genes should be pervasive. Here, we explore its effect on SA genetic variation in fitness using a two locus model with negative epistasis. Our results demonstrate that epistasis often increases the parameter space showing polymorphism for SA loci. This is because selection in one locus is affected by allele frequencies at the other, which can act to balance net selection in males and females. Increased linkage between SA loci had more marginal effects. We also show that under some conditions, large portions of the parameter space evolve to a state where male benefit alleles are fixed at one locus and female benefit alleles at the other. This novel effect of epistasis on SA loci, which we term the ‘equity effect’, may have important effects on population differentiation and may contribute to speciation. More generally, these results support the suggestion that epistasis contributes to population divergence.